Pattern and dynamics of the ground vegetation in south Swedish Carpinus betulus forests: importance of soil chemistry and management

The vegetation and environmental conditions of south Swedish hornbeam Carpinus betulus forests are described with data from 35 permanent sample plots The main floristic gradient of the ground vegetation is closely related to acid-base properties of the top soil base saturation, pH and organic matter content Other floristic differences are related to tree canopy cover and the distance of the sample plots to the Baltic coast Species richness of herbaceous plants typical of forests increases with soil pH, The number of other herbaceous species, occurring in both forests and open habitats, and of woody species is not related to pH Comparisons of vegetation data from 1983 and 1993 show relatively small compositional differences of the herbaceous forest flora The number of other herbaceous species increased considerably m those plots where canopy trees had been cut after 1983 The number of new species in managed plots increases with soil pH Species losses and gains of the herbaceous forest flora between 1983 and 1993 are generally lower as compared with other herbaceous species and woody species However, the ground cover of herbaceous forest species, especially of Oxalis acetosella and Lamium galeobdolon , was considerably lower in 1993 as compared to 1983 in both unmanaged and managed plots Possible explanations for this decrease are current soil acidification and drought during the growing season     

Abundance of trees and grasses in a woodland savanna in relation to environmental factors

The relationships of plant species associations and underlying environmental factors in a woodland savanna in South Africa were investigated. 40 plots were included with 25 tree and 17 grass species dominating the arboreal and ground cover. Correspondence Analysis described the relationships between soil moisture retention, soil nutrients and the abundance of trees and grasses. Dry matter indices represented the accumulated effects of rainfall, fire and grazing of the herbaceous layer. Variations in the abundance of plants corresponded to well-defined gradients of soil nutrients. The distribution of grass and tree species along the ordination axes indicated that soils with high water retention capacity and high nutrient contents provided a suitable substrate for many of the tree species sampled. However, grass species abundance was high in plots with porous soils and poor nutrient availability.     

Patterns of Fine Root Mass and Distribution along a Disturbance Gradient in a Tropical Montane Forest, Central Sulawesi (Indonesia)

Large parts of the remaining tropical moist forests of South-east Asia are encroached at their margins by selective logging, rattan harvesting and the establishment of small agroforest plantations under the rainforest canopy. These slight to heavy disturbances affect aboveground forest structure by reducing wood biomass and canopy cover; however, they may also have a profound impact on the belowground compartment. In a lower montane moist forest of Central Sulawesi, we studied the profile totals of fine root biomass (FRB_tot, roots <2 mm until 50 cm of soil depth) and of fine root necromass (FRN_tot), the vertical distribution of fine root mass, and the fine root live/dead ratio by root coring in 12 forest stands that represented a gradient in forest use (or disturbance) intensity (forest use type A: undisturbed natural forest, B and C: slightly or moderately disturbed forests with selective timber extraction, D: heavily disturbed cacao agroforest systems under a remaining rainforest cover; each forest types being replicated three times). FRB_tot decreased significantly from forest A to the disturbed B, C and D forests, and reached less than 60% of the FRB_tot value of A in the agroforest systems D. A similar decrease with increasing disturbance intensity was found for FRN_tot. Forest disturbance intensity had no significant influence on the vertical distribution of fine root biomass in the profiles. According to correlation and principal components analyses, fractional canopy cover was the most important factor influencing FRB_tot and FRN_tot, whereas diameter at breast height, stand basal area, stem density, soil pH and base saturation had only a minor or no influence on root mass. A reduction in canopy cover from 90% (forest type A) to 75% (types C and D) was associated with a reduction in FRB_tot by about 45% which indicates that timber extraction leads not only to canopy gaps but to corresponding ‘root gaps’ in the soil as well. We conclude that forest encroachment that is widespread in large parts of South-east Asia’s remaining rainforests significantly reduces tree fine root biomass and associated carbon sequestration, even if it is conducted at moderate intensities only.     

Development of Brachystegia‐Julbernardia woodland after clear‐felling in central Zambia: Evidence for high resilience

Question: Does the development of Brachystegia‐Julbernardia (miombo) woodland after felling, and under a variable fire regime, occur via a serai stage of fire‐tolerant species? Location: Four sites in central Zambia, Africa. Methods: Trees in replicate plots were clear‐cut and stumps and resprouts enumerated. Species recruited into the tree layer (> 2.0 m tall) were monitored for 11 years (1991–2001) and fire occurrence and herbaceous biomass assessed annually to determine fuel loads. Results: Fire frequency was variable at the study sites and fuel loads were generally too low to suppress woodland regeneration after felling. However, at one site a change from low to high fire frequency arrested woodland development and triggered a regression towards a ‘fire‐trap’ vegetation type in which a few fire‐tolerant species survived. There was no evidence to support the hypothesis that miombo woodland regeneration is facilitated by a sere of fire‐tolerant species. All regrowth after felling was from resprouting plants present before felling. Trees with a previous history of felling sprouted more vigorously than trees that had not been felled before. Species richness in the tree layer increased with time since felling because resprout species had different height growth rates. Conclusion: The resilience of miombo trees after clear‐felling is largely due to their capacity to regenerate vegetatively from resprouts and stumps after release from frequent fires. Coppicing is therefore recommended as a suitable management technique for miombo woodland in central southern Africa.     

Carbon storage,structure and composition of miombo woodlands in Tanzania’s Eastern Arc Mountains

We determine the aboveground biomass and carbon storage (ABGC) of trees and the herbaceous layer in miombo woodland in the Eastern Arc Mountains (EAM) of Tanzania. In four 1‐ha sample plots in Nyanganje and Kitonga Forests, we measured all trees ≥10 cm diameter alongside height and wood mass density. The plots contained an average of 20 tree species ha^?1 (range 11–29) and 344 stems ha^?1 (range 281–382) with Shannon diversity values of 1.05 and 1.25, respectively. We weighted nine previously published woody savannah allometric models based on whether: (i) the model was derived from the same geographical region; (ii) the model included tree height/wood mass density in addition to stem diameter; and (iii) sample size was used to fit the model. The weighted mean ABGC storage from the nine models range from 13.5 ± 2 to 29.8 ± 5 Mg ha^?1. Measured ABGC storage in the herbaceous layer, using the wet combustion method, adds 0.55 ± 0.02 Mg C ha^?1. Estimates suggest that EAM miombo woodlands store a range of 13–30 Mg ha^?1 of carbon. Although the estimates suggest that miombo woodlands store significant quantities of carbon, caution is required as this is the first estimate based on in situ data.     

Overstorey Control of Understorey Species Composition in a Near-natural Temperate Broadleaved Forest in Denmark

Little is known about the importance of the forest overstorey relative to other factors in controlling the spatial variability in understorey species composition in near-natural temperate broadleaved forests. We addressed this question for the 19 ha ancient forest Suserup Skov (55°22′ N, 11°34′ E) in Denmark, one of the few old-growth temperate broadleaved forest remnants in north-western Europe, by inventorying understorey species composition and environmental conditions in 163 100 m² plots. We use unconstrained and constrained ordinations, variation partitioning, and Indicator Species Analysis to provide a quantitative assessment of the importance of the forest overstorey in controlling understorey species composition. Comparison of the gradients extracted by unconstrained and constrained ordinations showed that the main gradients in understorey species composition in our old-growth temperate broadleaved forest remnant are not caused by variability in the forest overstorey, but are related to topography and soil, edge effects, and unknown broad-scale factors. Nevertheless, overstorey-related variables uniquely accounted for 15% of the total explained variation in understorey species composition, with the pure overstorey-related (R_po), topography and soil (R_pt), edge and anthropogenic disturbance effects (R_pa), and spatial (R_ps) variation fractions being of equal magnitude. The forward variable selection showed that among the overstorey-related variables understorey light availability and to a lesser extent vertical forest structure were most important for understorey species composition. No unique influence of overstorey tree species identity could be documented. There were many indicator species for high understorey light levels and canopy gap centres, but none for medium or low light or closed canopy. Hence, no understorey species behaved as obligate shade plants. Our study shows that, the forest overstorey has a weak control of understorey species composition in near-natural broadleaved forest, in contrast to results from natural and managed forests comprising both conifer and broadleaved species. Nevertheless, >20% of the understorey species found were indicators of high light conditions or canopy openings. Hence, variability in canopy structure and understorey light availability is important for maintaining understorey species diversity.     

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r ²=0.71) than to the normalized difference vegetation index (NDVI, r ²=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.     

Response of floodplain understorey species to environmental gradients and tree invasion: a functional trait perspective

Plants are connected to habitats by functional traits which are filtered by environmental gradients. Since tree species composition in the forest canopy can influence ecosystem processes by changing resource availability, litter accumulation, and soil nutrient content, we hypothesised that non-native invasive trees can establish new environmental filters on the understorey communities. In the hardwood floodplain forests in Northern Italy, the invasive trees Robinia pseudoacacia L. and Prunus serotina Ehrh. are the dominant canopy species. We used trait data assembled from databases and iterative RLQ analysis to identify a parsimonious set of functional traits responding to environmental variables (soil, light availability, disturbance, and stand structure) and the dominant native and invasive canopy species. Then, RLQ and fourth-corner analysis was conducted to investigate the joint structure between macro-environmental variables and species traits and functional groups were identified. The trait composition of the herb-layer was significantly related to the main environmental gradients and the presence of the invaders in the canopy showed significant relationships with several traits. In particular, the presence of P. serotina may mitigate or even erase the effect of disturbances, maintaining a stable forest microclimate and thus favouring ‘true’ forest species, while R. pseudoacacia may slow down forest succession and regeneration by establishing new stable associations with a graminoid-dominated understorey. The impact of the two invasive trees on herb layer composition appears to differ, indicating that different management and control strategies may be needed.     

Effects of Wild Pig Disturbance on Forest Vegetation and Soils

In North America, wild pigs (Sus scrofa; feral pigs, feral swine, wild boars) are a widespread exotic species capable of creating large-scale biotic and abiotic landscape perturbations. Quantification of wild pig environmental effects has been particularly problematic in northern climates, where they occur only recently as localized populations at low densities. Between 2016 and 2017, we assessed short-term (within ~2 yrs of disturbance) effects of a low-density wild pig population on forest features in the central Lower Peninsula of Michigan, USA. We identified 16 8-ha sites using global positioning system locations from 7 radio-collared wild pigs for sampling. Within each site, we conducted fine-scale assessments at 81 plots and quantified potential disturbance by wild pigs. We defined disturbance as exposure of overturned soil, often resulting from rooting behavior by wild pigs. We quantified ground cover of plants within paired 1-m² frames at each plot, determined effects to tree regeneration using point-centered quarter sampling, and collected soil cores from each plot. We observed less percent ground cover of native herbaceous plants and lower species diversity, particularly for plants with a coefficient of conservatism ≥5, in plots disturbed by wild pigs. We did not observe an increase in colonization of exotic plants following disturbance, though the observed prevalence of exotic plants was low. Wild pigs did not select for tree species when rooting, and we did not detect any differences in regeneration of light- and heavy-seeded tree species between disturbed or undisturbed plots. Magnesium and ammonium content in soils were lower in disturbed plots, suggesting soil disturbance accelerated leaching of macronutrients, potentially altering nitrogen transformation. Our study suggested that disturbances by wild pigs, even at low densities, alters short-term native herbaceous plant diversity and soil chemistry. Thus, small-scale exclusion of wild pigs from vulnerable and rare plant communities may be warranted. © 2020 The Wildlife Society.     

Disturbance indicators and population decline of logged species in Mt. Elgon Forest,Kenya

Mount (Mt) Elgon forest in western Kenya is important for biodiversity, environmental protection and socio‐economic development. Characterizing forest conditions is essential for evaluation of sustainable management and conservation activities. This paper covers findings of a study which determined and analysed indicators useful in monitoring disturbance levels in the Mt Elgon Forest. A systematic survey was carried out and covered 305 plots of 0.02 ha and 250 smaller nested regeneration plots along 10 belt transects that were distributed in five blocks within the moist lower montane forest type. Collected and analysed data include types of disturbance, tree species composition, abundance and logged species. Correlation breakdown among disturbance types revealed that, paths were indicators of the number of tree harvesting sites (r_s =1.00, P < 0.01) and of de‐vegetated areas through grass harvesting (r_s = 0.90, P = 0.04). Solanum mauritianum Scop. was an indicator of old‐charcoal production sites. Logging targeted 13 tree species and harvested trees with diameter at breast height above 20 cm. The most exploited species were Olea capensis L. and Deinbolia kilimandscharica Taub. All exploited species had low regeneration but tree regeneration was not an effective indicator of logging.     

Leaf traits and associated ecosystem characteristics across subtropical and timberline forests in the Gongga Mountains,Eastern Tibetan Plateau

Knowledge of how leaf characteristics might be used to deduce information on ecosystem functioning and how this scaling task could be done is limited. In this study, we present field data for leaf lifespan, specific leaf area (SLA) and mass and area-based leaf nitrogen concentrations (N_mass, N_area) of dominant tree species and the associated stand foliage N-pool, leaf area index (LAI), root biomass, aboveground biomass, net primary productivity (NPP) and soil available-N content in six undisturbed forest plots along subtropical to timberline gradients on the eastern slope of the Gongga Mountains. We developed a methodology to calculate the whole-canopy mean leaf traits to include all tree species (groups) in each of the six plots through a series of weighted averages scaled up from leaf-level measurements. These defined whole-canopy mean leaf traits were equivalent to the traits of a leaf in regard to their interrelationships and altitudinal trends, but were more useful for large-scale pattern analysis of ecosystem structure and function. The whole-canopy mean leaf lifespan and leaf N_mass mainly showed significant relationships with stand foliage N-pool, NPP, LAI and root biomass. In general, as elevation increased, the whole-canopy mean leaf lifespan and leaf N_area and stand LAI and foliage N-pool increased to their maximum, whereas the whole-canopy mean SLA and leaf N_mass and stand NPP and root biomass decreased from their maximum. The whole-canopy mean leaf lifespan and stand foliage N-pool both converged towards threshold-like logistic relationships with annual mean temperature and soil available-N variables. Our results are further supported by additional literature data in the Americas and eastern China.     

山坡退化林地林木生长对微地形人工干预的响应

为探究水土保持微地形人工干预措施(等高反坡阶)对坡面退化林地林木生长的影响,在昆明市松华坝水源区内选取典型山坡退化林地,通过动态观测和对比分析不同样地林木胸径、树高、新枝生长量及样地叶面积指数(LAI)的变化差异,再结合土壤水分有效性分析以探讨差异原因.结果表明:等高反坡阶样地云南松胸径和树高的最大值和均值均大于对照样地(CK),退化林群落中小径阶和低矮的植株占更大比例,表明等高反坡阶样地中优势木新生植株和退化严重植株的生长潜力得到激发,不同样地云南松植株新枝平均枝长和枝径的年变化差异率达到了72.4%和39.1%.等高反坡阶样地云南松植株新枝生长量和样地LAI的变化及新枝生长的速度大于CK.土壤含水率和新枝生长量与LAI均呈显著相关,对水分有效性分析后得到等高反坡阶样地易效水比例(64.2%)整体高于CK(54.7%),5月至9月雨季中,等高反坡阶样地易效水存在时间更长,这有利于林木的生长,进而改变退化林地植物群落结构.     

Species composition and diversity of small Afromontane forest fragments in northern Ethiopia

In the highlands of northern Ethiopia, remnants of the original Afromontane forest vegetation are largely restricted to church yards and other sacred groves in a matrix of cropland and semiarid degraded savanna. To assess the potential for natural forest regeneration, species composition and diversity of all forest fragments (10) in a study area of 13,000 ha were analyzed in relation to environmental and soil variables. Using a random design and a density of approximately one plot per 2 ha in all fragments, 31 20×20 m plots were sampled. Indicator species analysis and MRPP tests yielded five communities representing two forest types and one degraded savanna habitat. The forest fragments had a species-poor tree and shrub community in which plots were rather homogeneous and most species abundant. NMDS and analysis of variance indicated that a topographical gradient correlated to soil phosphorus, soil depth, stoniness and the proximity to the river system explained the major differences in species composition and separated moist and dry Afromontane forest communities. The grazing intensity further partitioned the habitats. Present communities and their environmental correlates indicate that the secondary climax forest in the area probably consisted of dry Afromontane forest interlaced by broad strips of moist Afromontane forest along rivers and streams and not a continuous, mono-dominant Juniperus forest as is often presumed. Negative effects of the degraded matrix on forest fragments increased with decreasing patch area and increasing shape irregularity. Nevertheless, all remaining fragments are important for their role in the landscape ecology of the region as refuges and species pools and should be protected and managed accordingly. If seed dispersal from forest fragments into exclosures and subsequent tree recruitment are both successful, the vegetation type most likely to establish is Afromontane savanna woodland, and if managed properly, eventually dry Afromontane forest may arise. Increasing the size of small patches and placing forest plantations and exclosures in the vicinity of small forest fragments is expected to yield the most immediate results. This approach may increase the likelihood of patch colonisation by frugivorous forest birds and thus foster the regeneration of native woody species.     

Differences in Spider Community Composition among Adjacent Sites during Initial Stages of Oak Woodland Restoration

Fire suppression has altered the uplands of northern Mississippi (U.S.A.). Once blanketed by open oak woodlands, this region is now experiencing mesophytic tree invasion, canopy closure, reduced oak regeneration, and herbaceous understory loss. In an attempt to reestablish historical conditions, experimental restoration was initiated through thinning and burning treatments. Our study, part of a comprehensive monitoring effort, is the first to examine the impact of oak woodland restoration on the spider community and associated habitat structure. Samples measuring a variety of environmental variables and utilizing an array of spider collecting techniques were taken within four habitats located at the restoration site: fire‐suppressed forest, moderately treated forest, intensely treated forest, and old field. Two main conclusions resulted from this study. (1) Open‐habitat specialists responded positively to increased canopy openness regardless of the availability of herbaceous vegetation. (2) Woodland restoration increased spider diversity, perhaps through the formation of diverse habitat structure and/or by altering species dominance patterns. A rise in open‐habitat specialist diversity was observed as treatment intensity increased, with no compensatory reduction in the diversity of forest specialists. What remains to be seen is whether the continued transition to open woodland habitat will result in losses of forest specialist species. More aggressive overstory tree thinning is currently being administered to encourage the growth of herbaceous grasses and forbs, which will permit future tests of a hypothesized decline in forest specialists.     

Disturbance of the herbaceous layer after invasion of an eutrophic temperate forest by wild boar

Disturbances of the soil and the tree canopy are crucial factors determining the diversity, composition and biomass of the herbaceous layer in forests. This study presents a detailed account of ground vegetation in permanent plots surveyed before and after invasion of wild boar (Sus scrofa) to a temperate deciduous broadleaf forest. Specifically, we aimed to quantify the effect of wild boar rooting on cover, richness and composition of spring ephemerals, summer green herbs and saplings of woody species in relation to tree canopy cover. Rooting frequency in sample plots increased from 0% in 2010 to 61% in 2013. In heavily rooted plots, the mean cover of spring ephemeral geophytes (mainly Anemone nemorosa, A. ranunculoides and Ranunculus ficaria) decreased from 75% to 39% between 2010 and 2013. Species richness of summer green herbs generally increased between 2010 and 2013 and was additionally positively affected by heavy rooting and low canopy cover. Rooting also caused heterogenization of the herbaceous layer and amplified ongoing compositional changes induced by changing light conditions. Frequency and richness of spring ephemeral and woody species remained unchanged. We conclude that overall species richness of the herbaceous layer may increase in the short‐term as a result of increased plant recruitment and seed dispersal. However, wild boar rooting can greatly reduce the ground cover of spring ephemerals in eutrophic broadleaf forests, thereby threatening their important ecological function. To avoid long‐term losses of characteristic spring flora elements, local population control of wild boar is necessary to reduce abundance and frequency of soil rooting.     

Severe dry winter affects plant phenology and carbon balance of a cork oak woodland understorey

Mediterranean climates are prone to a great variation in yearly precipitation. The effects on ecosystem will depend on the severity and timing of droughts. In this study we questioned how an extreme dry winter affects the carbon flux in the understorey of a cork oak woodland? What is the seasonal contribution of understorey vegetation to ecosystem productivity?We used closed-system portable chambers to measure CO₂ exchange of the dominant shrub species (Cistus salviifolius, Cistus crispus and Ulex airensis), of the herbaceous layer and on bare soil in a cork oak woodland in central Portugal during the dry winter year of 2012. Shoot growth, leaf shedding, flower and fruit setting, above and belowground plant biomass were measured as well as seasonal leaf water potential. Eddy-covariance and micrometeorological data together with CO₂ exchange measurements were used to access the understorey species contribution to ecosystem gross primary productivity (GPP).The herbaceous layer productivity was severely affected by the dry winter, with half of the yearly maximum aboveground biomass in comparison with the 6 years site average. The semi-deciduous and evergreen shrubs showed desynchronized phenophases and lagged carbon uptake maxima. Whereas shallow-root shrubs exhibited opportunistic characteristics in exploiting the understorey light and water resources, deep rooted shrubs showed better water status but considerably lower assimilation rates. The contribution of understorey vegetation to ecosystem GPP was lower during summer with 14% and maximum during late spring, concomitantly with the lowest tree productivity due to tree canopy renewal. The herbaceous vegetation contribution to ecosystem GPP never exceeded 6% during this dry year stressing its sensitivity to winter and spring precipitation.Although shrubs are more resilient to precipitation variability when compared with the herbaceous vegetation, the contribution of the understorey vegetation to ecosystem GPP can be quite variable and will ultimately depend of tree density and canopy cover.     

Phylogenetic constraints to soil properties determine elevational diversity gradients of forest understory vegetation

Elevational diversity gradients (EDGs) of vegetation are shaped by the evolutionary histories of plants as well as by ecological factors. However, few studies of EDGs have focused on the roles of phylogenetic constraints and the effects of complicated interactions among environmental factors. Here, we examine the direct and indirect effects of environmental factors in forming EDGs of forest understory vegetation. The study plots were selected along elevational gradients in cool-temperate and sub-alpine forests in the University of Tokyo Chichibu Forest of central Japan. Tree seedlings and herbs were identified, and environmental factors (elevation, soil temperature, soil pH, soil CN ratio, forest type, basal area, canopy openness, and slope) were measured in these plots. Structural equation modeling (SEM) including taxonomic and phylogenetic diversity was used to consider the causal relationships between environmental conditions and the diversity of understory vegetation. In addition, phylogenetic signals in habitat requirements were tested. The taxonomic and phylogenetic diversities of tree seedlings increased monotonically with elevation, and the same pattern was found for the taxonomic diversity of herbs. The SEM indicated that both the taxonomic and phylogenetic diversity of tree seedlings were most affected by soil properties, although the phylogenetic diversity of herbs was determined by light conditions. These results highlight the importance of environmental filtering by soil properties in shaping EDGs of tree seedlings. This study implies that phylogenetic constraints in the adaptation to soil properties should be considered when predicting changes in EDGs under environmental fluctuations.     

Variation in leaf area index and stand leaf mass of European beech across gradients of soil acidity and precipitation

Leaf area index (LAI, the one-sided foliage area per unit ground surface area) is a key determinant of plant productivity which has a large influence on water and energy exchange between vegetation and the atmosphere. The variation in forest LAI across landscapes and environmental gradients and its causes are not sufficiently understood. We measured the LAI of European beech (Fagus sylvatica) by litter trapping in 23 closed, mature stands across gradients of rainfall and soil acidity or fertility. With a mean LAI of 7.4 m² m⁻² (minimum: 5.6, maximum: 9.5 m² m⁻²), beech stands maintained a comparably high leaf area index with relatively small variation along steep environmental gradients. Contrary to expectation, decreasing water availability (rainfall gradient from 1030 to 520 mm yr⁻¹) or increasing soil acidity (pH 3–7) had no significant effect on LAI. Stand leaf mass (M _l) increased slightly with soil fertility (C/N ratio, base saturation). We regressed parameters of site water availability (rainfall), soil fertility or acidity (pH, base saturation, C/N ratio, exchangeable Mg and Al content), and stand structure (stand age and stem density) against LAI and M _l in order to detect environmental controls of stand leaf area. Stand age was the most influential factor for both LAI and M _l (negative relationship). Stem density and the base saturation of the soil affected M _l significantly, but had a weak influence on LAI. We conclude that the leaf area index of beech is mainly under control of age-related physiological factors, whereas the influence of soil chemistry and rainfall is comparably low.     

Seedling regeneration,environment and management in a semi‐deciduous African tropical rain forest

Questions: How is seedling regeneration of woody species of semi‐deciduous rain forests affected by (a) historical management for combinations of logging, arboricide treatment or no treatment, (b) forest community type and (c) environmental gradients of topography, light and soil nutrients? Location: Budongo Forest Reserve, Uganda. Methods: Seedling regeneration patterns of trees and shrubs in relation to environmental factors and historical management types were studied using 32 0.5‐ha plots laid out in transects along a topographic gradient. We compared seedling species diversity, composition and distribution patterns along topographic gradients and within types of historical management regimes and forest communities to test whether environmental factors contributed to differences in species composition of seedlings. Results: A total of 85 624 woody seedlings representing 237 species and 46 families were recorded in this rain forest. Cynometra alexandri C.H. Wright and Lasiodiscus mildbraedii Engl. had high seedling densities and were widely distributed throughout the plots. The most species‐rich families were Euphorbiaceae, Fabaceae, Rubiaceae, Meliaceae, Moraceae and Rutaceae. Only total seedling density was significantly different between sites with different historical management, with densities highest in logged, intermediate in logged/arboricided and lowest in the nature reserve. Forest communities differed significantly in terms of seedling diversity and density. Seedling composition differed significantly between transects and forest communities, but not between topographic positions or historical management types. Both Chao‐Jaccard and Chao‐Sørensen abundance‐based similarity estimators were relatively high in the plot, forest community and in terms of historical management levels, corroborating the lack of significant differences in species richness within these groups. The measured environmental variables explained 59.4% of variance in seedling species distributions, with the three most important being soil organic matter, total soil titanium and leaf area index (LAI). Total seedling density was positively correlated with LAI. Differences in diversity of >2.0 cm dbh plants (juveniles and adults) also explained variations in seedling species diversity. Conclusions: The seedling bank is the major route for regeneration in this semi‐deciduous tropical rain forest, with the wide distribution of many species suggesting that these species regenerate continuously. Seedling diversity, density and distribution are largely a function of adult diversity, historical management type and environmental gradients in factors such as soil nutrient content and LAI. The species richness of seedlings was higher in soils both rich in titanium and with low exchangeable cations, as well as in logged areas that were more open and had a low LAI.     

Beneath the veil: plant growth form influences the strength of species richness–productivity relationships in forests

Aim Species richness has been observed to increase with productivity at large spatial scales, though the strength of this relationship varies among functional groups. In forests, canopy trees shade understorey plants, and for this reason we hypothesize that species richness of canopy trees will depend on macroclimate, while species richness of shorter growth forms will additionally be affected by shading from the canopy. In this study we test for differences in species richness–productivity relationships (SRPRs) among growth forms (canopy trees, shrubs, herbaceous species) in small forest plots. Location We analysed 231 plots ranging from 34.0° to 48.3° N latitude and from 75.0° to 124.2° W longitude in the United States. Methods We analysed data collected by the USDA Forest Inventory and Analysis program for plant species richness partitioned into different growth forms, in small plots. We used actual evapotranspiration as a macroclimatic estimate of regional productivity and calculated the area of light‐blocking tissue in the immediate area surrounding plots for an estimate of the intensity of local shading. We estimated and compared SRPRs for different partitions of the species richness dataset using generalized linear models and we incorporated the possible indirect effects of shading using a structural equation model. Results Canopy tree species richness increased strongly with regional productivity, while local shading primarily explained the variation in herbaceous plant richness. Shrub species richness was related to both regional productivity and local shading. Main conclusions The relationship between total forest plant species richness and productivity at large scales belies strong effects of local interactions. Counter to the pattern for overall richness, we found that understorey herbaceous plant species richness does not respond to regional productivity gradients, and instead is strongly influenced by canopy density, while shrub species richness is under multivariate control.